A comparative study of the functional response of four species of carnivorous stoneflies

SUMMARY 1. Comparisons were made of the functional responses of mature larvae of Perlodes microcephalus, Isoperla grammatica, Dinocras cephalotes and Perla bipunctata. Experiments were performed in stream tanks with natural substrata and glass bottoms, so that feeding could be observed above and below the substratum. There was one stonefly per tank and one of 10 prey densities between 20 and 200 larvae of either Chironomus sp. or Baetis rhodani per tank. Consumed prey were replaced in a first set of experiments but not in a second set. Additional experiments assessed intraspecific interference between larvae of each predator species (two to five predators per tank). 2. The number of prey eaten increased curvilinearly with prey density. The relationship was described by two models, a Type II instantaneous model and its integrated equivalent, for experiments with and without prey replacement, respectively. Handling time did not change significantly with prey density, and was the same for experiments with and without prey replacement. Estimates of attack rate were similar for the two models, but varied between prey type and predator species. Handling time varied considerably but was normally distributed for each prey type and predator species. Mean handling time varied for Chironomus from 39 s for Isoperla, which rarely ate a whole larva, to 57 s for Perlodes and for Baetis from 116 s for Perlodes to 167 s for Perla. All predators were more efficient at capturing Baetis, but the longer handling time for Baetis ensured that more Chironomus were eaten. It was concluded that these variations in attack rate and handling time were related to activity and growth differences between the predator species, and that experiments with and without prey replacement could both be relevant to the field, depending on how the predators searched for food. 3. In the interference experiments, mean handling time did not change with increasing predator density, but attack rate decreased curvilinearly, the decrease varying from negligible for Isoperla to marked for Perlodes. Prey capture decreased with decreasing attack rate. Therefore, interference reduced prey consumption, but this effect was negligible for Isoperla and increasingly severe in the order Dinocras, Perla and Perlodes.     

Prey switching in four species of carnivorous stoneflies

1. Previous studies compared the functional responses to their prey, and both intraspecific and interspecific interference, in mature larvae of Dinocras cephalotes, Perla bipunctata, Isoperla grammatica and Perlodes microcephalus. The present study examines switching by larvae of the same species presented with different proportions of two contrasting prey types; larvae of Baetis rhodani and Chironomus sp. In each experiment, 200 prey were arranged in nine different combinations of the two prey types (20 : 180, 40 : 160, 60 : 140, 80 : 120, 100 : 100, 120 : 80, 140 : 60, 160 : 40, 180 : 20). Prey were replaced as they were eaten. A model predicted the functional response in the absence of switching and provided a null hypothesis against which any tendency for switching could be tested. 2. No evidence for prey switching by Dinocras and Perla was obtained, both species showing a slight preference for Baetis over Chironomus. Prey switching occurred in Isoperla and Perlodes. As the relative abundance of one prey type increased in relation to the alternative, the proportion eaten of the former prey changed from less to more than expected from its availability, the relationship being described by an S‐shaped curve. Isoperla and Perlodes switched to a preference for Baetis when its percentage of the total available prey exceeded 57 and 42%, respectively. Equivalent values for Chironomus were 43 and 58% for Isoperla and Perlodes, respectively. Switching was strongest in Perlodes. 3. Non‐switching in Dinocras and Perla was related to their feeding strategy, both species being more successful when using a non‐selective ambush strategy at dusk and dawn rather than a search strategy during the night. Both Isoperla and Perlodes used a search strategy. The smaller Isoperla fed chiefly at dusk and dawn, and preferred Chironomus larvae, whereas most of the larger Perlodes fed continuously from dusk to dawn and preferred Baetis larvae.     

Contrasting diel activity and feeding patterns of four species of carnivorous stoneflies

1. The hypothesis under test was that larvae of Dinocras cephalotes (Curtis), Perla bipunctata Pictet, Isoperla grammatica (Poda), and Perlodes microcephalus (Pictet) differed markedly in their diel activity and feeding patterns. Mature larvae collected about 1 month prior to adult emergence were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks with natural substrata and glass bottoms, so that activity could be observed above and below the substratum. A dull red light was used for observations in the dark. Food (larvae of Ephemeroptera, Simuliidae, and Chironomidae) was provided in excess. 2. Larvae for gut analyses were taken 1 h before dusk or dawn (n = 30 larvae per species for each day or night sample). The only species with food in the gut for the day samples was P. microcephalus. All species fed at night, the mean number of prey per larva being very similar for D. cephalotes, P. bipunctata, and I. grammatica but significantly higher for P. microcephalus. Most prey were insect larvae, especially Simuliidae and Chironomidae. 3. Diel activity patterns of single larvae differed interspecifically but not intraspecifically. Larvae of D. cephalotes and P. bipunctata were rarely active during the day; their activity increased at dusk and decreased at dawn, and was highest during the night; their success at prey capture was highest at dusk and dawn, with an ambush rather than a search strategy. Isoperla grammatica was rarely active during the day, most active at dusk and dawn when prey capture was highest, using a search strategy, and less active for the rest of the night. Perlodes microcephalus was active during the day, but only below the substratum, and very active from dusk to dawn with a high prey‐capture success, using a search strategy. These experiments provided support for the hypothesis under test. The discussion concludes that the results could also help to explain known differences in growth rate and the length of the life‐cycle in these four species.     

Predator density and competition modify the benefits of group formation in a shoaling reef fish

Synthesis Predation risk experienced by individuals living in groups depends on the balance between predator dilution, competition for refuges, and predator interference or synergy. These interactions operate between prey species as well: the benefits of group living decline in the presence of an alternative prey species. We apply a novel model‐fitting approach to data from field experiments to distinguish among competing hypotheses about shifts in predator foraging behavior across a range of predator and prey densities. Our study provides novel analytical tools for analyzing predator foraging behavior and offers insight into the processes driving the dynamics of coral reef fish. Studies of predator foraging behavior typically focus on single prey species and fixed predator densities, ignoring the potential importance of complexities such as predator dilution; predator‐mediated effects of alternative prey; heterospecific competition; or predator–predator interactions. Neglecting the effects of prey density is particularly problematic for prey species that live in mixed species groups, where the beneficial effects of predator dilution may swamp the negative effects of heterospecific competition. Here we use field experiments to investigate how the mortality rates of a shoaling coral reef fish (a wrasse: Thalassoma amblycephalum), change as a result of variation in: 1) conspecific density, 2) density of a predator (a hawkfish: Paracirrhites arcatus), and 3) presence of an alternative prey species that competes for space (a damselfish: Pomacentrus pavo). We quantify changes in prey mortality rates from the predator's perspective, examining the effects of added predators or a second prey species on the predator's functional response. Our analysis highlights a model‐fitting approach that discriminates amongst multiple hypotheses about predator foraging in a community context. Wrasse mortality decreased with increasing conspecific density (i.e. mortality was inversely density‐dependent). The addition of a second predator doubled prey mortality rates, without significantly changing attack rate or handling time – i.e. there was no evidence for predator interference. The presence of a second prey species increased wrasse mortality by 95%; we attribute this increase either to short‐term apparent competition (predator aggregation) or to a decrease in handling time of the predator (e.g. through decreased wrasse vigilance). In this system, 1) prey benefit from intraspecific group living though a reduced predation risk, and 2) the benefit of group living is reduced in the presence of an alternative prey species.     

Temperature modulates the effects of predation and competition on mosquito larvae

1. Predation risk affects interspecific competition by decreasing foraging activity and relative competitive ability. Predation risk is determined by predators' prey choice and prey responses, both of which can be influenced by temperature. Temperature is especially important for larval prey and can result in a trade‐off between predator‐induced decreases in foraging activity and growth. Interspecific competition must also be examined in relation to intraspecific density‐dependent competition; weaker interspecific competition leads to coexistence of competitors. 2. This study explored how temperature (15 and 25 °C) could affect a focal species, larvae of the mosquito Culex quinquefasciatus, by examining prey choice in a shared predator (mosquitofish; Gambusia holbrooki) and the effects of predation risk on interspecific competition with Limnodynastes peronii tadpoles. Intraspecific density‐dependent competition in C. quinquefasciatus at these temperatures was also examined. 3. At 25 °C, G. holbrooki consumption of both C. quinquefasciatus and L. peronii increased; however, the effects of interspecific competition on mosquito survival did not decrease with L. peronii exposure to predation risk. The relationship between intraspecific density‐dependent competition and interspecific competition was temperature‐dependent, with competitive dominance of L. peronii at 25 °C. Male and female mosquitoes had different temperature‐dependent responses, indicating sex‐specific intrinsic responses to starvation and differential selection pressures. At 25 °C, females were susceptible to interspecific competition by L. peronii, while males were susceptible to intraspecific competition. 4. The use of competitors as biological controls has implications for mosquito disease transmission, and these results suggest that control effectiveness may be modified by climate change.     

The diet of two species of Isoperla (Plecoptera: Perlodidae) in relation to season,site, and sympatry

The seasonal change in gut contents of nymphs of Isoperla grammatica and I. difformis from six streams in southern Sweden was analysed. Both species had ingested a variety of benthic prey and vegetable matter, predominantly diatoms. Some seasonality was evident with high percentages of diatoms in spring in I. grammatica, and in autumn in I. difformis. The scope of food was larger in the latter species which contained about equal proportions of vegetable matter, chironomids, mayfly, stonefly, and black fly larvae. In I. grammatica plant matter and chironomids dominated strongly, comprising > 50% of the gut contents on an annual basis, > 90 % in spring. While small nymphs of I. difformis contained a low proportion of animal matter, only gradually increasing with size, the nymphs of I. grammatica were carnivorous from very early instars. Both species switched to a temporarily strong utilization of algae in spring. This differed among sites, and appeared to reflect differences in insolation and thus the availability of algae. There was a significant negative relationship between the mean densities of Isoperla nymphs and the proportion of animal material found in the guts of I. grammatica (R ² = 0.86). Considering the density of I. grammatica alone, the relationship was weaker (R ² = 0.56). A positive correlation between predator and prey size was observed. With chironomid prey the size range increased with predator size. With simuliid prey, however, prey size increased with predator size in such a way that it suggests selection rather than just an expanding prey size range. Correlations were stronger and regression coefficients significantly higher for I. grammatica than for I. difformis. We suggest that I. grammatica, which ingests a much wider size range of prey might choose prey of optimal sizes more readily than the more synchronously developing I. difformis. Although the life cycles of the two species are staggered, overlap in size distribution indicates that competition for food could be important in spring. However, observed differences in diet should facilitate coexistence. Gut content differences might in turn be accomplished through microhabitat segregation.     

The microdistribution of some lotic insect predators in relation to their prey and to abiotic factors

SUMMARY. 1. The microdistribution of three species of insect predators in a Swedish stream was assessed using a multivariate statistical approach. Both abiotic factors and factors pertaining to the prey community were included in the analysis.
2. The factors most strongly correlated to the distribution of large Dinocras cephalotes were the densities of case less caddis larvae ( Wormaldia sp. and Hydropsyche siltalai ) and the median weight of stonefly prey. For small D. cephalotes , the density of H. siltalai and the percentage of moss cover were most important. Isoperla grammatica showed a significant correlation to blackfly density. Rhyacophila nubila occurred predominantly in patches with high prey biomass and with high densities and median weight of case less caddis larvae.
3. It was concluded that the quantity and quality of the prey, such as size, availability and species, influenced the predators'microdistribution. The distributional pattern differed between species of predators and was probably related to their different hunting strategies.     

Does Mutual Interference Affect the Feeding Rate of Aphidophagous Coccinellids? A Modeling Perspective

Mutual interference involves direct interactions between individuals of the same species that may alter their foraging success. Larvae of aphidophagous coccinellids typically stay within a patch during their lifetime, displaying remarkable aggregation to their prey. Thus, as larvae are exposed to each other, frequent encounters may affect their foraging success. A study was initiated in order to determine the effect of mutual interference in the coccinellids’ feeding rate. One to four 4^th larval instars of the fourteen-spotted ladybird beetle Propylea quatuordecimpunctata were exposed for 6 hours into plastic containers with different densities of the black bean aphid, Aphis fabae, on potted Vicia faba plants. The data were used to fit a purely prey-dependent Holling type II model and its alternatives which account for interference competition and have thus far been underutilized, i.e. the Beddington-DeAngelis, the Crowley-Martin and a modified Hassell-Varley model. The Crowley-Martin mechanistic model appeared to be slightly better among the competing models. The results showed that although the feeding rate became approximately independent of predator density at high prey density, some predator dependence in the coccinellid’s functional response was observed at the low prey—high predator density combination. It appears that at low prey densities, digestion breaks are negligible so that the predators do waste time interfering with each other, whereas at high prey densities time loss during digestion breaks may fully accommodate the cost of interference, so that the time cost may be negligible.     

Combined versus Single Species Release of Predaceous Mites: Predator–Predator Interactions and Pest Suppression

Interactions such as competition, intraguild predation (IGP), and cannibalism affect the development and coexistence of predator populations and can have significance for biological control of commonly exploited pest organisms. We studied the consequences of combined versus single release of two predaceous mite species (Phytoseiidae), with differing degrees of diet specialization, on their population dynamics and the suppression of the carmine spider mite, Tetranychus cinnabarinus Boisduval (Tetranychidae), on greenhouse-grown gerbera. Population growth of the specialist predator Phytoseiulus persimilis Athias-Henriot was greater and population decline steeper when released in combination with the generalist Neoseiulus californicus McGregor than when released alone. In contrast, the N. californicus population grew and declined more gradually when released in combination with P. persimilis, compared to the single species release. The differential impact on each other's population dynamics can be primarily attributed to contrasting properties in competition, IGP, and cannibalism. At the same overall predator density and as long as prey was abundant, the specialist P. persimilis was more strongly affected by intraspecific competition than by interspecific competition with the generalist N. californicus. In contrast, interspecific competition with P. persimilis had a greater impact on N. californicus than intraspecific competition. After prey depletion, the generalist predator N. californicus was more likely to engage in IGP than was the specialist predator P. persimilis. Overall, the study demonstrates that prey specificity has significance for the quality and intensity of predator–predator interactions and indicates potential implications for biological control of spider mites. All predator releases (i.e., either species alone and both species in combination) resulted in reduction of the spider mite population to zero density. Individual release of the specialist P. persimilis led to the most rapid spider mite suppression. Nonetheless, in perennial greenhouse-grown crops P. persimilis and N. californicus could have complementary effects and a combination of the two predators could enhance long-term biological control of spider mites. The potential risks and benefits associated with the release of both species are discussed.     

High intraguild predator density induces thinning effects on and increases temporal overlap with prey populations

Intraguild (IG) predator density can alter its effects on intraguild prey populations through several mechanisms, including density-dependent processes that affect IG predator traits such as size or growth that enhance or limit its predatory abilities. We examined whether intraspecific density-dependence altered IG predator traits, as well as the subsequent interspecific effects among its intraguild prey within a larval salamander guild. Four densities of ringed salamanders (Ambystoma annulatum), the IG predator, were combined with the presence/absence of spotted salamanders (A. maculatum), the IG prey, within experimental mesocosms. We modeled the effects of A. annulatum density on both conspecific and heterospecific responses that would be indicative of density-dependent competition and predation, respectively. We also modeled the reciprocal interspecific effects of A. maculatum on A. annulatum. We found that increasing intraspecific density negatively affected morphological traits but not survival of A. annulatum. No interspecific effects of A. maculatum on A. annulatum were observed. Alternatively, traits of A. maculatum showed nonlinear relationships with increasing A. annulatum density. Thinning effects of A. annulatum on A. maculatum were observed, as survival was positively and size negatively related for A. maculatum with IG predator density. The temporal overlap of the IG predator and prey also increased nonlinearly with IG predator density, intensifying the potential encounter rate of the two species. Overall, this study shows that density-dependent processes in IG predators can significantly affect traits of both themselves, as well as IG prey, which could ultimately change whether competition or predation occurs between the two groups.     

Intraspecific competition in immature Amblyseius fallacis,Amblyseius andersoni,Typhlodromus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

Intraspecific competition in immature Amblyseius fallacis, Amblyseius andersoni, Typhlodromus occidentalis and Typhlodromus pyri was examined in the laboratory using small cages at five different predator densities (two, four, eight, 16 and 32) in the absence and presence of prey 100 eggs of two-spotted spider mite, Tetranychus urticae (Koch), at 25 ± 1°C, 80% RH and 16L:8D photoperiod. In the absence of spider mite prey, some individuals of immature phytoseiids showed increased development and surival with increasing predator densities up to certain limits, but none survived to the adult stage, except for a single male each of A. andersoni and A. fallacis who completed development by cannibalizing on conspecifics at a density of 32 predators per cage. In the absence of spider mite prey, the mean immature survival time was independent of the initial predator density, but the variance of survival time increased with predator density. In the presence of prey, the proportion of immatures surviving to adulthood generally decreased with initial predator density and dropped sharply to almost none at the predator density of 32 for A. fallacis, eight for A. andersoni, 16 for T. occidentalis and four for T. pyri. The number of prey consumed per predator during the first day generally decreased with predator density in all four species, as prey available per predator decreased and the competition for food increased with predator density. Our data indicate that scramble competition is operating in these four species. Although cannibalism was occasionally observed, especially after the exhaustion of prey and in the generalist predators such as A. andersoni, the immatures of these phytoseiids were less influenced by the interference of conspecifics than by the increasing difficulty of finding food at high predator densities. The implications of this study for understanding phytoseiid population dynamics and their use in biological control are discussed.     

Intra‐ and interspecific interactions between aphidophagous ladybirds: the role of prey in predator coexistence

Cannibalism (CANN) and intraguild predation (IGP) may provide energy and nutrients to individuals and eliminate potential competitors. These negative competitive interactions could also affect the coexistence of predatory species. The co‐occurrence of aphidophagous ladybird species in crops creates opportunities for CANN and IGP, especially when aphids become scarce. The Lotka–Volterra model predicts the coexistence of two species if intraspecific competition is stronger than interspecific interference interactions. Cycloneda sanguinea L. and Eriopis connexa (Germar) (both Coleoptera: Coccinellidae) coexist in sweet pepper crops in La Plata (Argentina) consuming mainly Myzus persicae (Sulzer) (Hemiptera: Aphididae). The present study used laboratory experiments to estimate levels of CANN and IGP by adults and larvae on eggs, and by adults on larvae, in both the presence and absence of prey (i.e., M. persicae), to explain the effect of prey on coexistence of these two predators. Levels of CANN by C. sanguinea and E. connexa were high in the absence of aphids, and decreased when prey was present. Intraguild predation was bidirectional and asymmetric. Adults and larvae of E. connexa were more voracious IG predators of C. sanguinea than vice versa, the former being the stronger IG predator and interference competitor. Eriopis connexa always won when larvae of the same instar were compared, whereas the larger larva always won when larvae were of different instars, regardless of species. In the presence of prey, CANN by both species decreased, but IGP by E. connexa on C. sanguinea remained high, suggesting that E. connexa could displace C. sanguinea via interspecific interference competition. Other factors potentially affecting the coexistence of C. sanguinea and E. connexa in sweet pepper crops are discussed.     

Nymphal feeding habits of Perla madritensis Rambur 1842 (Plecoptera: Perlidae)

The diet of Perla madritensis, endemic species of the northern half of the Iberian Peninsula, is described for the first time by means of a study carried out in north-western Spain. As other species of the genus Perla, this taxon behaves mainly as predator, with Chironomidae, followed by Baetidae, being the most abundant prey in its gut. Shifts in diet composition were detected in relation to size in this species, with smaller prey being replaced by larger ones such as Simuliidae and Leptophlebiidae when the nymphs become larger.     

Influence of predator density on nonindependent effects of multiple predator species

Interactions between multiple predator species are frequent in natural communities and can have important implications for shared prey survival. Predator density may be an important component of these interactions between predator species, as the frequency of interactions between species is largely determined by species density. Here we experimentally examine the importance of predator density for interactions between predator species and subsequent impacts on prey. We show that aggressive interactions between the predatory shore crabs Carcinus maenas and Hemigrapsus sanguineus increased with predator density, yet did not increase as fast as negative interactions between conspecifics. At low density, interactions between conspecific and heterospecific predators had similar inhibitory impacts on predator function, whereas conspecific interference was greater than interference from heterospecifics at high predator density. Thus the impact of conspecific interference at high predator density was sufficient in itself that interactions with a second predator species had no additional impact on per capita predation. Spatial and temporal variability in predator density is a ubiquitous characteristic of natural systems that should be considered in studies of multiple predator species.     

Competitive interactions among two specialist predators and a generalist predator of hemlock woolly adelgid, Adelges tsugae (Hemiptera: Adelgidae) in south-western Virginia

1 Competitive interactions among two specialist predators, Laricobius nigrinus and Sasajiscymnus (Pseudoscymnus) tsugae, and a generalist predator, Harmonia axyridis Pallas, of hemlock woolly adelgid, Adelges tsugae were evaluated in hemlock stands in south‐western Virginia. The two specialist predators are part of a biological control program for A. tsugae, and the potential for competition among these species and previously established generalist predators in the field is unknown. 2 Adult predators were evaluated in branch cages during spring and summer at two field sites infested with A. tsugae. Using females only in 2003 and sexual pairs in 2004, predator survival and net reproduction were examined, as well as their feeding and impact on A. tsugae when present alone and in conspecific and heterospecific groupings. 3 Predator survival was not affected by the presence of additional predators. Total feeding was greater for all species when placed in predator groupings, suggesting that interactions do not significantly interfere with feeding activity. Net reproduction per predator was negatively affected by conspecifics, but unaffected by heterospecifics, indicating that direct or indirect intraspecific interference may occur. In spring, L. nigrinus showed the greatest impact on A. tsugae, and H. axyridis had the greatest impact during summer. 4 These results suggest that it would be beneficial to utilize multiple predator species combinations over single species when implementing biological control for A. tsugae. Low‐density releases are also recommended to reduce intraspecific interference.     

Prey: predator ratio dependence in the functional response of a freshwater amphipod

1. First known for their shredding activity, freshwater amphipods also behave as active predators with consequences for prey population regulation and amphipod coexistence in the context of biological invasions. 2. A way to quantify predation is to determine the average consumption rate per predator, also known as its functional response (FR). 3. Although amphipods are gregarious and can display social interactions that can alter per capita consumption rates, previous studies using the FR approach to investigate amphipod predation ignored such potential mutual interference because they did not consider variations in predator density. 4. We investigated the FR of Echinogammarus berilloni feeding on dipteran larvae with joint variations in prey and predator densities. This bivariate experimental design allowed us to estimate interference and to compare the fits of the three main classes of theoretical FR models, in which the predation rate is a function of prey density alone (prey‐dependent models), of both prey and predator densities (predator‐dependent models) or of the prey‐to‐predator ratio (ratio‐dependent models). 5. The Arditi–Ginzburg ratio‐dependent FR model provided the best representation of the FR of E. berilloni, whose predation rate showed a decelerating rise to a horizontal asymptote as prey abundance increased. 6. Ratio dependence means that mutual interference between amphipods leads to prey sharing. Mutual interference is likely to vary between amphipod species, depending on their level of aggressiveness.     

Notes on Plecoptera (19) Isoperla lugens (Klapalek, 1923), replaces I. alpicola Brinck, 1949

A male lectotype and several paralectotypes of Chloroperla lugens Klapálek, 1923 are designated. The species is transferred to genus Isoperla and is the valid name for Isoperla alpicola Brinck, 1949.     

Effect of the cyclopoid copepod Mesocyclops thermocyclopoides on the interactions between the predatory rotifer Asplanchna intermedia and its prey Brachionus calyciflorus and B. angularis

In many shallow, eutrophic subtropical ponds, brachionid rotifers are common prey of the predatory copepod Mesocyclops thermocyclopoides. The predatory rotifer Asplanchna intermedia, which is itself a potential prey of the copepod, also feeds preferentially on brachionids. We studied in the laboratory the population dynamics of two mutually competing prey species, Brachionus angularis and B. calyciflorus, in the presence of the two predators A. intermedia and M. thermocyclopoides. The experimental design included separate population dynamics studies with one prey–one predator, two prey–one predator, one prey–two predator, and two prey–two predator systems. These combinations were compared with controls, in which both the prey species (B. angularis and B. calyciflorus) were grown separately and in combination with each other. In the absence of any predator, B. angularis generally eliminated the larger B. calyciflorus. Selective predation by the copepod allowed B. calyciflorus to persist longer in competition with B. angularis. Feeding by M. thermocyclopoides on A. intermedia reduced the predation pressure on B. calyciflorus. However, given enough time, the cyclopoid copepod was able to eliminate both the brachionids as well as the predatory Asplanchna.     

Ontogenetic shifts in the functional response and interference interactions of Rhyacophila dorsalis larvae (Trichoptera)

1. Ontogenetic shifts in predator behaviour can affect the assessment of food‐web structure and the development of predator–prey models. Therefore, it is important to establish if the functional response and interference interactions differ between life‐stages. These hypotheses were tested by (i) comparing the functional response of second, third, fourth and fifth larval instars of Rhyacophila dorsalis, using three stream tanks with one Rhyacophila larva per tank and one of 10 prey densities between 20 and 200 larvae of Chironomus sp.; (ii) using other experiments to assess interference within instars (two to five larvae of the same instar per tank), and between pairs of different instars (one, two or three larvae per instar; total predator densities of two, four or six larvae per tank). 2. The first hypothesis was supported. The number of prey eaten by each instar increased with prey density, the relationship being described by a type II model. The curvilinear response was stronger for fourth and fifth instars than for second and third instars. Mean handling time did not change significantly with prey density, and increased with decreasing instar number from 169 s for fifth instars to 200 s for second instars. Attack rate decreased progressively with decreasing instar number. Handling time varied considerably for each predator–prey encounter, but was normally distributed for each predator instar. Variations in attack rate and handling time were related to differences in activity between instars, fourth and fifth instars being more active and aggressive than second and third instars, and having a higher food intake. 3. The second hypothesis was partially supported. In the interference experiments between larvae of the same instar or different instars, mean handling time did not change significantly with increasing predator density, and attack rate did not change for second and third instars but decreased curvilinearly for fourth and fifth instars. Interference between some instars could not be studied because insufficient second instars were available at the same time as fourth and fifth instars, and most third instars were eaten by fourth and fifth instars in the experiments. Prey capture always decreased with decreasing attack rate. Therefore, interference reduced prey consumption in fourth and fifth instars, but not in second and third instars. The varying feeding responses of different instars should be taken into account when assessing their role in predator–prey relationships in the field.     

Density of an intraguild predator mediates feeding group size, intraguild egg predation, and intra- and interspecific competition

Intraguild predation (IGP) is common in most communities, but many aspects of density-dependent interactions of IG predators with IG prey are poorly resolved. Here, we examine how the density of an IG predator can affect feeding group size, IG egg predation, and the growth responses of IG prey. We used laboratory feeding trials and outdoor mesocosm experiments to study interactions between a social intraguild predator (larvae of the wood frog; Rana sylvatica) and its prey (spotted salamander; Ambystoma maculatum). Larvae of R. sylvatica could potentially affect A. maculatum by consuming shared larval food resources or by consuming eggs and hatchlings. However, successful egg predation requires group feeding by schooling tadpoles. We established from five to 1,190 hatchlings of R. sylvatica in mesocosms, then added either 20 A. maculatum hatchlings to study interspecific competition, or a single egg mass to examine IGP. Crowding strongly suppressed the growth of R. sylvatica, and IGP was restricted to the egg stage. In the larval competition experiment, growth of A. maculatum was inversely proportional to R. sylvatica density. In the predation experiment, embryonic mortality of A. maculatum was directly proportional to the initial density of R. sylvatica and the mean number of tadpoles foraging on egg masses. IGP on eggs reduced A. maculatum hatchling density, which accelerated larval growth. Surprisingly, the density of R. sylvatica had no overall effect on A. maculatum growth because release from intraspecific competition via egg predation was balanced by increased interspecific competition. Our results demonstrate that the density of a social IG predator can strongly influence the nature and intensity of interactions with a second guild member by simultaneously altering the intensity of IGP and intra- and interspecific competition.L . A. Burley and A. T. Moyer contributed equally to this work.